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Abstract Isoprene, a volatile hydrocarbon, is typically emitted from the leaves of many plant species. Given its well‐known function in plant growth and defense aboveground, we examined its effects on root physiology. We used isoprene‐emitting (IE) lines and a non‐emitting (NE) line of Arabidopsis and investigated their performance by analyzing root phenotype, hormone levels, transcriptome, and metabolite profiles under both normal and salt stress conditions. We show that IE lines emitted tiny amounts of isoprene from roots and showed an increased root/shoot ratio compared with NE line. Isoprene emission exerted a noteworthy influence on hormone profiles related to plant growth and stress response, promoting root development and salt‐stress resistance. Methyl erythritol 4‐phosphate pathway metabolites, precursors of isoprene and hormones, were higher in the roots of IE lines than in the NE line. Transcriptome data indicated that the presence of isoprene increased the expression of key genes involved in hormone metabolism/signaling. Our findings reveal that constitutive root isoprene emission sustains root growth under saline conditions by regulating and/or priming hormone biosynthesis and signaling mechanisms and expression of key genes relevant to salt stress defense.

Plants are central to complex networks of multitrophic interactions. Increasing evidence suggests that beneficial microorganisms (BMs) may be used as plant biostimulants and pest biocontrol agents. We investigated whether tomato (Solanum lycopersicum) plants are thoroughly colonized by the endophytic and entomopathogenic fungus Beauveria bassiana, and how such colonization affects physiological parameters and the phenotype of plants grown under unstressed conditions or exposed to the pathogenic fungus Botrytis cinerea. As a positive control, a strain of the well-known biocontrol agent and growth inducer Trichoderma afroharzianum was used. As multitrophic interactions are often driven by (or have consequences on) volatile organic compounds (VOCs) released by plants constitutively or after induction by abiotic or biotic stresses, VOC emissions were also studied. Both B. bassiana and T. afroharzianum induced a significant but transient (one to two-day-long) reduction of stomatal conductance, which may indicate rapid activation of defensive (rejection) responses, but also limited photosynthesis. At later stages, our results demonstrated a successful and complete plant colonization by B. bassiana, which induced higher photosynthesis and lower respiration rates, improved growth of roots, stems, leaves, earlier flowering, higher number of fruits and yield in tomato plants. Beauveria bassiana also helped tomato plants fight B. cinerea, whose symptoms in leaves were almost entirely relieved with respect to control plants. Less VOCs were emitted when plants were colonized by B. bassiana or infected by B. cinerea, alone or in combination, suggesting no activation of VOC-dependent defensive mechanisms in response to both fungi.

ABSTRACTIn recent years, anthropogenic activities and climate change have significantly increased exposure of plants to environmental stresses (single or multiple) and pollutants, which negatively affect plant growth, survival, and productivity. Plants may activate an armament of defenses against such environmental stresses. Isoprene, the most abundant biogenic volatile organic compound emitted by plants, is supposed to induce stress tolerance directly, by quenching reactive oxygen species, or indirectly by strengthening photosynthetic membranes and reprogramming expression of genes that are involved in antioxidant defense mechanisms. On the other hand, isoprene is also involved in tropospheric chemistry that contributes to the production of air pollutants when mixing with anthropogenic gases. In this review, we summarized current knowledge about the impact of air and soil pollutants on isoprene emission from plants, focusing on possible feedback and feedforward mechanisms that may affect whole ecosystem functioning and evolution of plant species. Despite limited available information, especially about long-term effects of soil pollutants, it may be speculated that isoprene generally improves fitness of plants challenged by air and soil pollutants, and their interaction with other organisms.

The increase in soil salinization represents a current challenge for plant productivity, as most plants, including crops, are mainly salt-sensitive species. The identification of molecular traits underpinning salt tolerance represents a primary goal for breeding programs. In this scenario, the study of intraspecific variability represents a valid tool for investigating natural genetic resources evolved by plants in different environmental conditions. As a model system, Arabidopsis thaliana, including over 750 natural accessions, represents a species extensively studied at phenotypic, metabolic, and genomic levels under different environmental conditions. Two haplogroups showing opposite root architecture (shallow or deep roots) in response to auxin flux perturbation were identified and associated with EXO70A3 locus variations. Here, we studied the influence of these genetic backgrounds on plant salt tolerance. Eight accessions belonging to the two haplogroups were tested for salt sensitivity by exposing them to moderate (75 mM NaCl) or severe (150 mM NaCl) salt stress. Salt-tolerant accessions were found in both haplogroups, and all of them showed efficient ROS-scavenging ability. Even if an exclusive relation between salt tolerance and haplogroup membership was not observed, the modulation of root system architecture might also contribute to salt tolerance.

A growing population coupled with a higher demand for food is putting pressure on agriculture. The use of synthetic pesticides and chemical fertilizers allowed us to boost agricultural productions, but at a great environmental cost. Exploitation of beneficial microorganism (BM)-plant interactions has been proposed as an eco-friendly solution to improve plant resistance to stresses and to increase productivity sustainably. We provide an overview of scientific evidence that this positive interaction is often mediated also by the release of microbial Volatile Organic Compounds (mVOCs). A few mVOCs are reported to have a double, not mutually exclusive, positive effect on plants, as plant growth promoters, and/or inducers of resistance against biotic and abiotic stress factors. They may also alter plant VOCs indirectly improving plant performances. However, mechanisms and functions of mVOCs need deeper investigation. By understanding mVOC modes of action on plants, further tools for sustainably improving plant productivity in agro-ecosystems may become soon available.

Abstract The growing world population and global increases in the standard of living both result in an increasing demand for food, feed and other plant‐derived products. In the coming years, plant‐based research will be among the major drivers ensuring food security and the expansion of the bio‐based economy. Crop productivity is determined by several factors, including the available physical and agricultural resources, crop management, and the resource use efficiency, quality and intrinsic yield potential of the chosen crop. This review focuses on intrinsic yield potential, since understanding its determinants and their biological basis will allow to maximize the plant's potential in food and energy production. Yield potential is determined by a variety of complex traits that integrate strictly regulated processes and their underlying gene regulatory networks. Due to this inherent complexity, numerous potential targets have been identified that could be exploited to increase crop yield. These encompass diverse metabolic and physical processes at the cellular, organ and canopy level. We present an overview of some of the distinct biological processes considered to be crucial for yield determination that could further be exploited to improve future crop productivity.

Abstract One of the main impacts of climate change on agriculture production is the dramatic increase of saline (Na+) content in substrate, that will impair crop performance and productivity. Here we demonstrate how the application of smart technologies such as an in vivo sensor, termed bioristor, allows to continuously monitor in real-time the dynamic changes of ion concentration in the sap of Arundo donax L. (common name giant reed or giant cane), when exposed to a progressive salinity stress. Data collected in vivo by bioristor sensors inserted at two different heights into A. donax stems enabled us to detect the early phases of stress response upon increasing salinity. Indeed, the continuous time-series of data recorded by the bioristor returned a specific signal which correlated with Na+ content in leaves of Na-stressed plants, opening a new perspective for its application as a tool for in vivo plant phenotyping and selection of genotypes more suitable for the exploitation of saline soils.

Distinct photosynthetic physiologies are found within the Moricandia genus, both C3-type and C2-type representatives being known. As C2-physiology is an adaptation to drier environments, a study of physiology, biochemistry and transcriptomics was conducted to investigate whether plants with C2-physiology are more tolerant of low water availability and recover better from drought. Our data on Moricandia moricandioides (Mmo, C3), M. arvensis (Mav, C2) and M. suffruticosa (Msu, C2) show that C3 and C2-type Moricandias are metabolically distinct under all conditions tested (well-watered, severe drought, early drought recovery). Photosynthetic activity was found to be largely dependent upon the stomatal opening. The C2-type M. arvensis was able to secure 25–50% of photosynthesis under severe drought as compared to the C3-type M. moricandioides. Nevertheless, the C2-physiology does not seem to play a central role in M. arvensis drought responses and drought recovery. Instead, our biochemical data indicated metabolic differences in carbon and redox-related metabolism under the examined conditions. The cell wall dynamics and glucosinolate metabolism regulations were found to be major discriminators between M. arvensis and M. moricandioides at the transcription level.

Isoprene-emitting plants are better protected against thermal and oxidative stresses, which is a desirable trait in a climate-changing (drier and warmer) world. Here we compared the ecophysiological performances of transgenic isoprene-emitting and wild-type non-emitting tobacco plants during water stress and after re-watering in actual environmental conditions (400 ppm of CO2 and 28 °C of average daily temperature) and in a future climate scenario (600 ppm of CO2 and 32 °C of average daily temperature). Furthermore, we intended to complement the present knowledge on the mechanisms involved in isoprene-induced resistance to water deficit stress by examining the proteome of transgenic isoprene-emitting and wild-type non-emitting tobacco plants during water stress and after re-watering in actual climate. Isoprene emitters maintained higher photosynthesis and electron transport rates under moderate stress in future climate conditions. However, physiological resistance to water stress in the isoprene-emitting plants was not as marked as expected in actual climate conditions, perhaps because the stress developed rapidly. In actual climate, isoprene emission capacity affected the tobacco proteomic profile, in particular by upregulating proteins associated with stress protection. Our results strengthen the hypothesis that isoprene biosynthesis is related to metabolic changes at the gene and protein levels involved in the activation of general stress defensive mechanisms of plants.

Volatile organic compounds (VOCs) from ‘emitting’ plants inform the ‘receiving’ (listening) plants of impending stresses or simply of their presence. However, the receptors that allow receivers to detect the volatile cue are elusive. Most likely, plants (as animals) have odorant-binding proteins (OBPs), and in fact, a few OBPs are known to bind ‘stress-induced’ plant VOCs. We investigated whether these and other putative OBPs may bind volatile constitutive and stress-induced isoprenoids, the most emitted plant VOCs, with well-established roles in plant communication and defense. Molecular docking simulation experiments suggest that structural features of a few plant proteins screened in databases could allow VOC binding. In particular, our results show that monoterpenes may bind the same plant proteins that were described to bind other stress-induced VOCs, while the constitutive hemiterpene isoprene is unlikely to bind any investigated putative OBP and may not have an info-chemical role. We conclude that, as for animal, there may be plant OBPs that bind multiple VOCs. Plant OBPs may play an important role in allowing plants to eavesdrop messages by neighboring plants, triggering defensive responses and communication with other organisms.

Arundo donax has been recognized as a promising crop for biomass production on marginal lands due to its superior productivity and stress tolerance. However, salt stress negatively impacts A. donax growth and photosynthesis. In this study, we tested whether the tolerance of A. donax to salinity stress can be enhanced by the addition of 5-aminolevulinic acid (ALA), a known promoter of plant growth and abiotic stress tolerance. Our results indicated that root exposure to ALA increased the ALA levels in leaves along the A. donax plant profile. ALA enhanced Na+ accumulation in the roots of salt-stressed plants and, at the same time, lowered Na+ concentration in leaves, while a reduced callose amount was found in the root tissue. ALA also improved the photosynthetic performance of salt-stressed apical leaves by stimulating stomatal opening and preventing an increase in the ratio between abscisic acid (ABA) and indol-3-acetic acid (IAA), without affecting leaf methanol emission and plant growth. Supply of ALA to the roots reduced isoprene fluxes from leaves of non-stressed plants, while it sustained isoprene fluxes along the profile of salt-stressed A. donax. Thus, ALA likely interacted with the methylerythritol 4-phosphate (MEP) pathway and modulate the synthesis of either ABA or isoprene under stressful conditions. Overall, our study highlights the effectiveness of ALA supply through soil fertirrigation in preserving the young apical developing leaves from the detrimental effects of salt stress, thus helping of A. donax to cope with salinity and favoring the recovery of the whole plant once the stress is removed.

MicroRNAs (miRNAs) are a class of non-coding molecules involved in the regulation of a variety of biological processes. They have been identified and characterized in several plant species, but only limited data are available for Arundo donax L., one of the most promising bioenergy crops. Here we identified, for the first time, A. donax conserved and novel miRNAs together with their targets, through a combined analysis of high-throughput sequencing of small RNAs, transcriptome and degradome data. A total of 134 conserved miRNAs, belonging to 45 families, and 27 novel miRNA candidates were identified, along with the corresponding primary and precursor miRNA sequences. A total of 96 targets, 69 for known miRNAs and 27 for novel miRNA candidates, were also identified by degradome analysis and selected slice sites were validated by 5′-RACE. The identified set of conserved and novel candidate miRNAs, together with their targets, extends our knowledge about miRNAs in monocots and pave the way to further investigations on miRNAs-mediated regulatory processes in A. donax, Poaceae and other bioenergy crops.

Isoprene is a small lipophilic molecule synthesized in plastids and abundantly released into the atmosphere. Isoprene-emitting plants are better protected against abiotic stresses, but the mechanism of action of isoprene is still under debate. In this study, we compared the physiological responses and proteomic profiles of Arabidopsis which express the isoprene synthase (ISPS) gene and emit isoprene with those of non-emitting plants under both drought-stress (DS) and well-watered (WW) conditions. We aimed to investigate whether isoprene-emitting plants displayed a different proteomic profile that is consistent with the metabolic changes already reported. Only ISPS DS plants were able to maintain the same photosynthesis and fresh weight of WW plants. LC–MS/MS-based proteomic analysis revealed changes in protein abundance that were dependent on the capacity for emitting isoprene in addition to those caused by the DS. The majority of the proteins changed in response to the interaction between DS and isoprene emission. These include proteins that are associated with the activation of secondary metabolisms leading to ABA, trehalose, and proline accumulations. Overall, our proteomic data suggest that isoprene exerts its protective mechanism at different levels: under drought stress, isoprene affects the abundance of chloroplast proteins, confirming a strong direct or indirect antioxidant action and also modulates signaling and hormone pathways, especially those controlling ABA synthesis. Unexpectedly, isoprene also alters membrane trafficking.

Isoprene is a highly reactive biogenic volatile hydrocarbon that strongly influences atmospheric oxidation chemistry and secondary organic aerosol budget. Many phytoplanktons emit isoprene like terrestrial pants. Planktonic isoprene emission is stimulated by light and heat and is seemingly dependent on photosynthesis, as in higher plants. However, prominent isoprene-emitting phytoplanktons are known to survive also as mixotrophs and heterotrophs. Chlorella vulgaris strain G-120, a unicellular green alga capable of both photoautotrophic and heterotrophic growth, was examined for isoprene emission using GC-MS and real-time PTR-MS in light (+CO2) and in darkness (+glucose). Chlorella emitted isoprene at the same rate both as a photoautotroph under light, and as an exclusive heterotroph while feeding on exogenous glucose in complete darkness. By implication, isoprene synthesis in eukaryotic phytoplankton can be fully supported by glycolytic pathways in absence of photosynthesis, which is not the case in higher plants. Isoprene emission by chlorophyll-depleted mixotrophs and heterotrophs in darkness serves unknown functions and may contribute to anomalies in oceanic isoprene estimates.

The methylerythritol 4-phosphate (MEP) pathway of isoprenoid biosynthesis produces chlorophyll side chains and compounds that function in resistance to abiotic stresses, including carotenoids, and isoprene. Thus we investigated the effects of moderate and severe drought on MEP pathway function in the conifer Picea glauca, a boreal species at risk under global warming trends. Although moderate drought treatment reduced the photosynthetic rate by over 70%, metabolic flux through the MEP pathway was reduced by only 37%. The activity of the putative rate-limiting step, 1-deoxy-D-xylulose-5-phosphate synthase (DXS), was also reduced by about 50%, supporting the key role of this enzyme in regulating pathway metabolic flux. However, under severe drought, as flux declined below detectable levels, DXS activity showed no significant decrease, indicating a much-reduced role in controlling flux under these conditions. Both MEP pathway intermediates and the MEP pathway product isoprene incorporate administered 13CO2 to high levels (75–85%) under well-watered control conditions indicating a close connection to photosynthesis. However, this incorporation declined precipitously under drought, demonstrating exploitation of alternative carbon sources. Despite the reductions in MEP pathway flux and intermediate pools, there was no detectable decline in most major MEP pathway products under drought (except for violaxanthin under moderate and severe stress and isoprene under severe stress) suggesting that the pathway is somehow buffered against this stress. The resilience of the MEP pathway under drought may be a consequence of the importance of the metabolites formed under these conditions.

It is well-acknowledged that plants in urban areas provide multiple ecosystem services. They contribute improving ambient quality and mitigating negative impacts of human presence, beautifying the anthropic environment, and promoting place identity and cultural heritage. However, the existence of plants in general, and trees in particular, cannot be considered independent on urban activities and infrastructures. Release of plant volatile compounds is profoundly affected in urban environments, in turn modifying plant relationships with other living organisms, both plants and animals, and affecting air chemistry and quality. Plants also interfere with stone artifacts, cultural and historical heritage. Plant-human coexistence requires precise and adequate managing measures, which have often been ignored in cities' government and planning. Plants and humans (and human infrastructures) are frequently considered as independent from each other and plant requirements are often disregarded, thus causing difficult or erroneous management and/or environmental damage. We review some of the most important ecosystem services provided by plants in urban environment, and also focus on possible negative effects of plants that may become relevant if urban vegetation is improperly managed and unintegrated in proper city planning, both of historical centers and of new towns or suburbs.

Environmental degradation and the decrease of ecosystem service provision are currently of major concern, with current agricultural systems being a major driver. To meet our future environmental and sustainability targets a transformation of the agro-food systems and current agricultural value chain are crucial. One approach to redesign farming systems is the concept of biodiversity-based agriculture (BBA) which relies on sustainable diversification of biological components and their natural interactions in farming systems to maximize fertility, productivity, and resilience to external perturbations. Despite minimizing anthropogenic inputs, BBA is not yet able to meet all beneficial environmental objectives. BBA applied in the Mediterranean basin requires urgent innovation in approaches, methodologies, and models for small-holder traditional farming systems to ensure a stable provision of ecosystem services and better resilience to environmental stresses linked to climate change. Legumes are the backbone of the Mediterranean agro-ecosystems from ancient times, but their unique and wide biodiversity was not sufficiently valorized, especially by North-African countries. Here, we present LEGU-MED, a three-year international project funded by PRIMA initiative 2019. An international consortium was established involving five universities, 5 research institutes, and one private company from 8 countries: Italy, Germany, Spain, Algeria, Tunisia, Turkey, Lebanon, and Croatia. The main objective of this project is to put forward an international and well-integrated plan to valorize the legume agrobiodiversity of the Mediterranean in biodiversity-based farming systems and consequently enhance agro-ecosystem functions and services in the Mediterranean basin. The successful completion of LEGU-MED will have the following impacts on Mediterranean legume-based farming systems: (1) improve water use efficiency, (2) reduce the use of anthropogenic inputs through the maintenance of soil fertility, (3) enhance pollination and improve ecological connectivity with flora and fauna, (4) protect close-by wildland ecosystems, (5) enhance other ecosystem services (e.g., pest, disease, and weed suppression), and (6) provide healthier and safer protein-rich food.

The realization of the full objectives of international policies targeting global food security and climate change mitigation, including the United Nation’s Sustainable Development Goals, the Paris Climate Agreement COP21 and the European Green Deal, requires that we (i) sustainably increase the yield, nutritional quality and biodiversity of major crop species, (ii) select climate-ready crops that are adapted to future weather dynamic and (iii) increase the resource use efficiency of crops for sustainably preserving natural resources. Ultimately, the grand challenge to be met by agriculture is to sustainably provide access to sufficient, nutritious and diverse food to a worldwide growing population, and to support the circular bio-based economy. Future-proofing our crops is an urgent issue and a challenging goal, involving a diversity of crop species in differing agricultural regimes and under multiple environmental drivers, providing versatile crop-breeding solutions within wider socio-economic-ecological systems. This goal can only be realized by a large-scale, international research cooperation. We call for international action and propose a pan-European research initiative, the CropBooster Program, to mobilize the European plant research community and interconnect it with the interdisciplinary expertise necessary to face the challenge.

The Mediterranean climate is characterized by hot dry summers and frequent droughts. Mediterranean crops are frequently subjected to high evapotranspiration demands, soil water deficits, high temperatures, and photo-oxidative stress. These conditions will become more severe due to global warming which poses major challenges to the sustainability of the agricultural sector in Mediterranean countries. Selection of crop varieties adapted to future climatic conditions and more tolerant to extreme climatic events is urgently required. Plant phenotyping is a crucial approach to address these challenges. High-throughput plant phenotyping (HTPP) helps to monitor the performance of improved genotypes and is one of the most effective strategies to improve the sustainability of agricultural production. In spite of the remarkable progress in basic knowledge and technology of plant phenotyping, there are still several practical, financial, and political constraints to implement HTPP approaches in field and controlled conditions across the Mediterranean. The European panorama of phenotyping is heterogeneous and integration of phenotyping data across different scales and translation of “phytotron research” to the field, and from model species to crops, remain major challenges. Moreover, solutions specifically tailored to Mediterranean agriculture (e.g., crops and environmental stresses) are in high demand, as the region is vulnerable to climate change and to desertification processes. The specific phenotyping requirements of Mediterranean crops have not yet been fully identified. The high cost of HTPP infrastructures is a major limiting factor, though the limited availability of skilled personnel may also impair its implementation in Mediterranean countries. We propose that the lack of suitable phenotyping infrastructures is hindering the development of new Mediterranean agricultural varieties and will negatively affect future competitiveness of the agricultural sector. We provide an overview of the heterogeneous panorama of phenotyping within Mediterranean countries, describing the state of the art of agricultural production, breeding initiatives, and phenotyping capabilities in five countries: Italy, Greece, Portugal, Spain, and Turkey. We characterize some of the main impediments for development of plant phenotyping in those countries and identify strategies to overcome barriers and maximize the benefits of phenotyping and modeling approaches to Mediterranean agriculture and related sustainability.

There is an urgent need for new sustainable solutions to support agriculture in facing current environmental challenges. In particular, intensification of productivity and food security needs require sustainable exploitation of natural resources and metabolites. Here, we bring the attention to the agronomic potential of volatile organic compounds (VOCs) emitted from leaves, as a natural and eco-friendly solution to defend plants from stresses and to enhance crop production. To date, application of VOCs is often limited to fight herbivores. Here we argue that potential applications of VOCs are much wider, as they can also protect from pathogens and environmental stresses. VOCs prime plant’s defense mechanisms for an enhanced resistance/tolerance to the upcoming stress, quench reactive oxygen species (ROS), have potent antimicrobial as well as allelopathic effects, and might be important in regulating plant growth, development, and senescence through interactions with plant hormones. Current limits and drawbacks that may hamper the use of VOCs in open field are analyzed, and solutions for a better exploitation of VOCs in future sustainable agriculture are envisioned.

Modern plant phenotyping, often using non-invasive technologies and digital technologies, is an emerging science and provides essential information on how genetics, epigenetics, environmental pressures, and crop management (farming) can guide selection toward productive plants suitable for their environment. Thus, phenotyping is at the forefront of future plant breeding. Bibliometric science mapping is a quantitative method that analyzes scientific publications throughout the terms present in their title, abstract, and keywords. The aim of this mapping exercise is to observe trends and identify research opportunities. This allows us to analyze the evolution of phenotyping research and to predict emerging topics of this discipline. A total of 1,827 scientific publications fitted our search method over the last 20 years. During the period 1997–2006, the total number of publications was only around 6.1%. The number of publications increased more steeply after 2010, boosted by the overcoming of technological bias and by a set of key developments at hard and software level (image analysis and data storage management, automation and robotics). Cluster analysis evidenced three main groups linked to genetics, physiology, and imaging. Mainly the model plant “Arabidopsis thaliana” and the crops “rice” and “triticum” species were investigated in the literature. The last two species were studied when addressing “plant breeding,” and “genomic selection.” However, currently the trend goes toward a higher diversity of phenotyped crops and research in the field. The application of plant phenotyping in the field is still under rapid development and this application has strong linkages with precision agriculture. EU co-authors were involved in 41.8% of the analyzed papers, followed by USA (15.4%), Australia (6.0%), and India (5.6%). Within the EU, coauthors were mainly affiliated in Germany (35.8%), France (23.7%), and United Kingdom (18.4%). Time seems right for new opportunities to incentivize research on more crops, in real field conditions, and to spread knowledge toward more countries, including emerging economies. Science mapping offers the possibility to get insights into a wide amount of bibliographic information, making them more manageable, attractive, and easy to serve science policy makers, stakeholders, and research managers.

Isoprene (C5H8) is a small lipophilic, volatile organic compound (VOC), synthesized in chloroplasts of plants through the photosynthesis-dependent 2-C-methyl-D-erythritol 4-phosphate (MEP) pathway. Isoprene-emitting plants are better protected against thermal and oxidative stresses but only about 20% of the terrestrial plants are able to synthesize isoprene. Many studies have been performed to understand the still elusive isoprene protective mechanism. Isoprene reacts with, and quenches, many harmful reactive oxygen species (ROS) like singlet oxygen (1O2). A role for isoprene as antioxidant, made possible by its reduced state and conjugated double bonds, has been often suggested, and sometimes demonstrated. However, as isoprene is present at very low concentrations compared to other molecules, its antioxidant role is still controversial. Here we review updated evidences on the function(s) of isoprene, and outline contrasting indications on whether isoprene is an antioxidant directly scavenging ROS, or a membrane strengthener, or a modulator of genomic, proteomic and metabolomic profiles (perhaps as a secondary effect of ROS removal) eventually leading to priming of antioxidant plant defenses, or a signal of stress for neighbor plants alike other VOCs, or a hormone-like molecule, controlling the metabolic flux of other hormones made by the MEP pathway, or acting itself as a growth and development hormone.

In the context of climatic change, more severe and long-lasting droughts will modify the fitness of plants, with potentially worse consequences on the relict trees. We have investigated the leaf phenotypic (anatomical, physiological and biochemical) plasticity in well-watered, drought-stressed and re-watered plants of two populations of Platanus orientalis, an endangered species in the west of the Mediterranean area. The two populations originated in contrasting climate (drier and warmer, Italy (IT) population; more humid and colder, Bulgaria (BG) population). The IT control plants had thicker leaves, enabling them to maintain higher leaf water content in the dry environment, and more spongy parenchyma, which could improve water conductivity of these plants and may result in easier CO2 diffusion than in BG plants. Control BG plants were also characterized by higher photorespiration and leaf antioxidants compared to IT plants. BG plants responded to drought with greater leaf thickness shrinkage. Drought also caused substantial reduction in photosynthetic parameters of both IT and BG plants. After re-watering, photosynthesis did not fully recover in either of the two populations. However, IT leaves became thicker, while photorespiration in BG plants further increased, perhaps indicating sustained activation of defensive mechanisms. Overall, our hypothesis, that plants with a fragmented habitat (i.e., the IT population) lose phenotypic plasticity but acquire traits allowing better resistance to the climate where they became adapted, remains confirmed.

The functional reasons for isoprene emission are still a matter of hot debate. It was hypothesized that isoprene biosynthesis evolved as an ancestral mechanism in plants adapted to high water availability, to cope with transient and recurrent oxidative stresses during their water-to-land transition. There is a tight association between isoprene emission and species hygrophily, suggesting that isoprene emission may be a favorable trait to cope with occasional exposure to stresses in mesic environments. The suite of morpho-anatomical traits does not allow a conservative water use in hygrophilic mesophytes challenged by the environmental pressures imposed or exacerbated by drought and heat stress. There is evidence that in stressed plants the biosynthesis of isoprene is uncoupled from photosynthesis. Because the biosynthesis of isoprene is costly, the great investment of carbon and energy into isoprene must have relevant functional reasons. Isoprene is effective in preserving the integrity of thylakoid membranes, not only through direct interaction with their lipid acyl chains, but also by up-regulating proteins associated with photosynthetic complexes and enhancing the biosynthesis of relevant membrane components, such as mono- and di-galactosyl-diacyl glycerols and unsaturated fatty acids. Isoprene may additionally protect photosynthetic membranes by scavenging reactive oxygen species. Here we explore the mode of actions and the potential significance of isoprene in the response of hygrophilic plants when challenged by severe stress conditions associated to rapid climate change in temperate climates, with special emphasis to the concomitant effect of drought and heat. We suggest that isoprene emission may be not a good estimate for its biosynthesis and concentration in severely droughted leaves, being the internal concentration of isoprene the important trait for stress protection.

The response of broad bean (Vicia faba) plants to water stress alone and in combination with green stink bug (Nezara viridula) infestation was investigated through measurement of: (1) leaf gas exchange; (2) plant hormone titres of abscisic acid (ABA) and its metabolites, and of salicylic acid (SA); and (3) hydrogen peroxide (H2O2) content. Furthermore, we evaluated the effects of experimentally water-stressed broad-bean plants on N. viridula performance in terms of adult host–plant preference, and nymph growth and survival. Water stress significantly reduced both photosynthesis (A) and stomatal conductance (gs), while infestation by the green stink bug had no effects on photosynthesis but significantly altered partitioning of ABA between roots and shoots. Leaf ABA was decreased and root ABA increased as a result of herbivore attack, under both well-watered and water-deprived conditions. Water stress significantly impacted on SA content in leaves, but not on H2O2. However, infestation of N. viridula greatly increased both SA and H2O2 contents in leaves and roots, which suggests that endogenous SA and H2O2 have roles in plant responses to herbivore infestation. No significant differences were seen for green stink bug choice between well-watered and water-stressed plants. However, for green stink bug nymphs, plant water stress promoted significantly lower weight increases and significantly higher mortality, which indicates that highly water-stressed host plants are less suitable for N. viridula infestation. In conclusion two important findings emerged: (i) association of water stress with herbivore infestation largely changes plant response in terms of phytohormone contents; but (ii) water stress does not affect the preference of the infesting insects, although their performance was impaired.

The function of dimethylsulphoniopropionate (DMSP) in plants is unclear. It has been proposed as an antioxidant, osmolyte and overflow for excess energy under stress conditions. The formation of DMSP is part of the methionine (MET) pathway that is involved in plant stress responses. We used a new analytical approach to accurately quantify the changes in DMSP concentration that occurred in two ecotypes of the biomass crop Arundo donax subject to moderate drought stress under field conditions. The ecotypes of A. donax were from a hot semi-arid habitat in Morocco and a warm-humid environment in Central Italy. The Moroccan ecotype showed more pronounced reductions in photosynthesis, stomatal conductance and photochemical electron transport than the Italian ecotype. An increase in isoprene emission occurred in both ecotypes alongside enhanced foliar concentrations of DMSP, indicative of a protective function of these two metabolites in the amelioration of the deleterious effects of excess energy and oxidative stress. This is consistent with the modification of carbon within the methyl-erythritol and MET pathways responsible for increased synthesis of isoprene and DMSP under moderate drought. The results of this study indicate that DMSP is an important adaptive component of the stress response regulated via the MET pathway in A. donax. DMSP is likely a multifunctional molecule playing a number of roles in the response of A. donax to reduced water availability.

Proton Transfer Reaction Mass Spectrometry (PTR-MS) has evolved in the last decade as a fast and high sensitivity sensor for the real-time monitoring of volatile compounds. Its applications range from environmental sciences to medical sciences, from food technology to bioprocess monitoring. Italian scientists and institutions participated from the very beginning in fundamental and applied research aiming at exploiting the potentialities of this technique and providing relevant methodological advances and new fundamental indications. In this review we describe this activity on the basis of the available literature. The Italian scientific community has been active mostly in food science and technology, plant physiology and environmental studies and also pioneered the applications of the recently released PTR-ToF-MS (Proton Transfer Reaction-Time of Flight-Mass Spectrometry) in food science and in plant physiology. In the very last years new results related to bioprocess monitoring and health science have been published as well. PTR-MS data analysis, particularly in the case of the ToF based version, and the application of advanced chemometrics and data mining are also aspects characterising the activity of the Italian community.

Dimethylsulphoniopropionate (DMSP) and dimethyl sulphide (DMS) are compounds found mainly in marine phytoplankton and in some halophytic plants. DMS is a globally important biogenic volatile in regulating of global sulphur cycle and planetary albedo, whereas DMSP is involved in the maintenance of plant-environment homeostasis. Plants emit minute amounts of DMS compared to marine phytoplankton and there is a need for hypersensitive analytic techniques to enable its quantification in plants. Solid Phase Micro Extraction from Head Space (HS-SPME) is a simple, rapid, solvent-free and cost-effective extraction mode, which can be easily hyphenated with GC-MS for the analysis of volatile organic compounds (VOCs). Using tomato (Solanum lycopersicum) plants subjected to water stress as a model system, we standardized a sensitive and accurate protocol for detecting and quantifying DMSP pool sizes, and potential DMS emissions, in cryoextracted leaves. The method relies on the determination of DMS free and from DMSP pools before and after the alkaline hydrolysis via HS-SPME-GC-MS. We found a significant (2.5 time) increase of DMSP content in water-stressed leaves reflecting clear stress to the photosynthetic apparatus. We hypothesize that increased DMSP, and in turn DMS, in water-stressed leaves are produced by carbon sources other than direct photosynthesis, and function to protect plants either osmotically or as antioxidants. Finally, our results suggest that SPME is a powerful and suitable technique for the detection and quantification of biogenic gases in trace amounts.

Most of the perennial plant species, particularly trees, emit volatile organic compounds (BVOCs) such as isoprene and monoterpenes, which in several cases have been demonstrated to protect against thermal shock and more generally against oxidative stress. In this paper, we show the response of three strong isoprene emitter species, namely, Phragmites australis, Populus x euramericana, and Salix phylicifolia exposed to artificial or natural warming of the root system in different conditions. This aspect has not been investigated so far while it is well known that warming the air around a plant stimulates considerably isoprene emission, as also shown in this paper. In the green house experiments where the warming corresponded with high stress conditions, as confirmed by higher activities of the main antioxidant enzymes, we found that isoprene uncoupled from photosynthesis at a certain stage of the warming treatment and that even when photosynthesis approached to zero isoprene emission was still ongoing. In the field experiment, in a typical cold-limited environment, warming did not affect isoprene emission whereas it increased significantly CO2 assimilation. Our findings suggest that the increase of isoprene could be a good marker of heat stress, whereas the decrease of isoprene a good marker of accelerated foliar senescence, two hypotheses that should be better investigated in the future.

Proton transfer reaction-time of flight (PTR-TOF) mass spectrometry was used to improve detection of biogenic volatiles organic compounds (BVOCs) induced by leaf wounding and darkening. PTR-TOF measurements unambiguously captured the kinetic of the large emissions of green leaf volatiles (GLVs) and acetaldehyde after wounding and darkening. GLVs emission correlated with the extent of wounding, thus confirming to be an excellent indicator of mechanical damage. Transient emissions of methanol, C5 compounds and isoprene from plant species that do not emit isoprene constitutively were also detected after wounding. In the strong isoprene-emitter Populus alba, light-dependent isoprene emission was sustained and even enhanced for hours after photosynthesis inhibition due to leaf cutting. Thus isoprene emission can uncouple from photosynthesis and may occur even after cutting leaves or branches, e.g., by agricultural practices or because of abiotic and biotic stresses. This observation may have important implications for assessments of isoprene sources and budget in the atmosphere, and consequences for tropospheric chemistry.